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The Seasonal Variation in the Cellulose Content of the Common Scottish Laminariaceae and Fucaceae

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The Seasonal Variation in the Cellulose Content of the Common Scottish Laminariaceae and Fucaceae 379 THE SEASONAL VARIATION IN THE CELLULOSE CONTENT OF THE COMMON SCOTTISH LAMINARIACEAE AND FUCACEAE By W. A. P. Black, B.Sc., Ph.D., F.R.I.C. Scottish Seaweed Research Association, Musselburgh (Text-figs. 1-6) INTRODUCTION Algal cellulose appears to have been relatively little studied and there is no information as to its seasonal variation in the brown marine algae. A study of the literature reveals numerous contradictions. In 1884, Stanford indicated in a report on alginic acid that he had obtained pure cellulose representing 10-15% of the air-dried plant. Kylin (1915) showed that algal cellulose gave the characteristic colour reaction with iodine and sulphuric acid. He boiled the material in turn with 1'25% sulphuric acid and 1'25% sodium hydroxide and called the insoluble residue cellulose. In 1926, Atsuki & Tomodo obtained a crude fibre figure of 6% for Lami- naria saccharina, but stated that the greater part of the crude fibre of the laminarias consisted of the hemicelluloses and that there was so far no evidence of the normal cellulose. Ricard (1931) removed the alginic acid with sodium carbonate, then treated the residue with dilute boiling solutions of sulphuric acid and potassium hydroxide. The hydrolysis of the residual material with sulphuric acid gave only traces of reducing sugars, and the author did not regard it as cellulose but called it algulose. L. jlexicaulis contained 4'3-7.6% of this material and L. saccharina2.8-10'9%. Dillon & O'Tuama (1935), however, isolated cellulose from the brown algae. The residue after the removal of alginic acid with dilute ammonia was treated with dilute hydrochloric acid and then several times with boiling 5% sodium hydroxide. It was finally washed with alcohol and ether, and was shown to have the properties of cellulose. On hydrolysis with hydrochloric acid it gave glucose characterizedby the isolation of the glucosazone,and with carbon disulphide it formed a viscose resembling the viscose from ordinary cellulose. They also prepared acetylated and methylated compounds with properties similar to those of the corresponding derivatives of ordinary cellulose. JOURN. MAR. BIOL. ASSOC. vol. XXIX. IQSO 25 380 W. A. P. BLACK Russel-Wells (I934) determined the crude fibre content of various algae and showed by its solubility in cuprammonium hydroxide and by its acetate that this fibre was cellulose. Viel (I939) carried out a systematic study of the celluloses from Fucus vesiculosus (2'IO %), F. serratus (2'81 %), Laminaria saccharina (6'9 %) and L. cloustoni (5'°4 %). Analysis showed the carbon-hydrogen percentages to be remarkably similar and in agreement with (C6H1o05)w Hydrolysis gave 90% glucose, while thermal fractionation of the products of pyrogenation gave graphs comparable with those of previously studied vegetable celluloses. Recent work by Percival & Ross (I949) has shown conclusively that algal cellulose is essentially the same as cotton cellulose. Hydrolysis with 72 % sulphuric acid gave only D-glucose. Cellobiose octa-acetate was prepared by acetolysis indicating the presence of I :4-,B-linkages. Determinations by periodate oxidations indicated a chain length of 160 units, but the original cellulose was no doubt degraded during isolation. Finally, X-ray diagrams of algal cellulose showed the characteristic pattern of normal cellulose. EXPERIMENTAL The samples analysed were those taken in 1946 and previously reported by the writer (I948a,b, 1949). The cellulose was estimated as follows. Ground seawe~d (2 g.) was boiled under reflux with sulphuric acid (200 m!., I'25 %) for 3° min., filtered by suction through a I x 2 Gallenkamp sintered glass crucible, washed free from acid with water, and the residue boiled with sodium hydroxide (200 m!., I'25%). Mter boiling in this way for 3° min., the solution was filtered through the I x 2 crucible and washed with water. The residue was removed from the crucible and kept overnight in chlorine water (IOOm!. saturated), filtered on a weighed hardened paper, washed free from chlorine and washed on the filter with hot sodium hydroxide solution (5° mI. N/lo). The residue was washed with water until free from alkali and finally with alcohol and ether, and then dried at 50° for IO min. to give a white product which was weighed. When applied to Fucus spp. the residue after treatment with boiling sodium hydroxide and chlorine water was often slimy and difficult to filter, and separa- tion by centrifuging was necessary. The results are given in Figs. 1-6. DISCUSSION OF RESULTS General Very little is known regarding the structure of the cell wall of the Phaeophy- ceae. Fritsch (I945) concludes that it consists of pectic substances with a layer giving the reactions of cellulose adjacent to the protoplast. In the brown algae the pectic substance is no doubt alginic acid which, with its long VARIATION IN CELLULOSE CONTENT 381 chain structure, will give flexibility to the plant. To withstand wave-action such a structure is probably reinforced with cellulose, and the results of this investigation show that the cellulose content increases with the depth of im- mersion, when additional strength is required by the plant. Very little is known, also, regarding the metabolism of the brown algae, but in the absence of any reducing sugars, it would appear that mannitol is probably the primary product of photosynthesis. In 1933, Khouvine showed that cellulose could be synthesized from mannitol by Acetobacter xylinum, and quite independently Barsha & Hibbert(1934) obtained the same results and showed that the cellulose membranes were chemically identical with cotton cellulose. In the brown algae, therefore, mannitol is probably the precursor of the cellulose. The Laminariaceae Laminaria cloustoni In Fig. I the graphs for the stipe, the frond and the whole plant show that the cellulose content on the anhydrous basis exhibits ~wo maxima and two minima in the year. Similar graphs have been obtained for several of the other constituents (Black, 1948a,b, 1949) and for the diatom periodicity, and some- times there is a correlation with the nutrient content of the water and the 'periods of rapid and slow growth (Black & Dewar, 1949). In March, when the new frond is forming, the cellulose is at a maximum of 5'76% in the frond and 10'27% in the stipe. From March to June/July a period of rapid growth occurs with an increase in the mannitol content. while the cellulose content decreases. This may be due to the accumulation of man- nitol in the whole plant or to elongation during rapid growth. From June to August/ September during the period of slow growth, when the nutrients are absent from the sea water, the cellulose content increases again, while at the same time laminarin is also increasing (Black, 1948a). In September, when the nutrients are again regenerated in the sea water, there is probably a second period of growth and a decrease in the cellulose content occurs. It would appear, therefore, that when the results are expressed on the anhydrous basis a correlation exists between the cellulose content and the periods of rapid and slow growth. When the results, however, are calculated on the wet basis (Fig. 2) two maxima are again recorded, one in February/March and the other from September to November. On the dry basis the cellulose content begins to fall in April and continues to fall until June; on the wet basis it is constant and at a minimum from April to June, due to the dry weight increasing with a rapid increase in mannitol. From June to November the cellulose on the wet basis increases while on the dry basis it begins to decrease in September, the increase in dry-weight content now being due to an increase in laminarin. A rapid decrease then occurs in December, the period of sporogenesis of this species, and this is probably accompanied by an increase in the uptake of water, as there is a considerable decrease in the dry-matter content both in 25-2 11 x~ x/.... X\ A 0 1'3 ; 1-2 x~ .~ '" 'Vi'8 .;;; .~ ~ 1'~ .J:I I " 1i::. ..>.. -0 3: cR~ 7- 4)'" 0 \ 8 'S /r-><\" /0 Qj 1<._X U ~ 6 I 0"0 51- " /°"-. c 0-70 . ° O 0, ~ / , / 0 " 0 ........ I I I I I <;).; / f . 41 2 3 4 5 6 7 8 9 10 11 12 ~~ 2 3 4 5 6 7 8 11 12 Month of year 1946 Monthof year 1946 . Fig. 1. Seasonal variation in the cellulose content of Laminaria cloustoni Fig. 2. Seasonal variation in the cellulose content of Laminaria cloustoni (dry basis): (A) in the stipes; (B) in the whole plant; (C) in the fronds. (wet basis): (A) in the stipes; (B) in the fronds. VARIATION IN CELLULOSE CONTENT 383 stipe and frond, which explains the rapid decrease on the wet basis while there is little change on the dry basis. On the wet basis, therefore, there would appear to be a correlation between the cellulose content and the reproductive cycle of the plant. Laminaria saccharina In this species the cellulose content (dry basis) of the frond (4-5%, Fig. 3) is of the same order of magnitude as that of L. cloustoni frond (Fig. I), but the stipes contain less cellulose (7-8%) than those of L. cloustoni (8'5-10%). Considerably less seasonal variation occurs in this species, but despite this the . two maxima are apparent in the year and occur.at approximately the same time as those of L. cloustoni. Laminaria digitata. The cellulose content of this species (Fig. 4) undergoes a similar seasonal variation to that of L. saccharina, with the exception of the loch frond samples in which the variation is between 3 and 5%.
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